BLUMEA 38 (1994) 313-330
Studies in Malesian Caesalpinioideae (Leguminosae). I.
The genera Acrocarpus, Afzelia, Copaifera, and Intsia
Ding Hou
Rijksherbarium / Hortus Botanicus, Leiden, The Netherlands
Summary
A partial, Malesian revision of the above listed four genera is given with special remarks on their
An Identification List is taxonomy, nomenclature, typification, relationship, distribution,etc. given.
Acrocarpus is so far known with only one species, A. fraxinifolius.
Afzelia has two widely distributed, variable species both occurring in Malesia: A. javanica and
A. rhomboidea.
Pseudosindora has been reduced to Copaifera, which consists of20-30 species mostly occurring in tropical America and Africa. One interesting species, C. palustris, is found in Malesia.
Intsia consists of two or more species. In Malesia there are two widely distributed, very variable,
difficult species, I. bijuga and I. palembanica. Information on their regeneration and uses have been reviewed.
Introduction
The Caesalpinioideae have been chosen by L. Watson and M.J. Dallwitz (Australia) for an automated data bank. They have prepared a comprehensive list of 134 charac- ters (Numbers 1-80 for general morphology). Their book, entitled 'The Genera of
Leguminosae-Caesalpinioideae, 1983', with descriptions and keys for 177 genera, has been produced by computer from that data bank. They have also noted under
characters which have been coded for lack of each genus those not yet (or recorded)
data. The descriptions are consistent and all characters are compared in parallel. For
and the reader is referred detailsconcerning their computer system programs used, to the Introduction and References of above mentionedbook.
of the Before the publication of that book, on my request, one authors, Dr Watson, sent me a detailed printout of the list of characters, keys, descriptions, etc. especially
cultivated in Malesia. In the of work- for those genera (incl. ones) occurring course
traditional this in Malesia, I have of those ing in a way on subfamily applied many
general morphological characters for species to facilitate using the DELTA system and associated programs which are now also available for MS-DOS microcomputers.
After having finished the descriptions I have mademodifications in order to con- form to a conventional format. The descriptions as presented here are as such in ac- cordance with the general generic treatment ofWatson and Dallwitz; they are far too in detailed for a Flora Malesiana treatment and seem superfluous especially case of small genera. 314 BLUMEA VOL. 38, No. 2, 1994
consist of tall Four small genera, viz. Acrocarpus, Afzelia, Copaifera, and Intsia,
(timber) tree species in Malesia. These trees resemble one another and are seldom
fruit. be without much in the field. in flower or They may recognized difficulty
Vegetative characters of the Malesian species of these genera have been described
in with for field rather more detail, together a separate paragraph compiled notes or recorded vernacular names, which may be helpful for identificationof sterile mate- rial.
Afzelia J.E. Sm. (Africa), Pahudia Miq. (Malesia), and Intsia Thouars (Madagas-
India car) are three closely alliedgenera. Prain (Sci. Mem.Med. Off. Army 12,1901,
of and Pa- 33-49, f. A-D) made a comprehensive study Afzelia its related genera
and hudia and Intsia on the generic delimitation, relationship, distribution, nomen-
Pahudia and Intsia is distinct clature. He concludedthatAfzelia and are congeneric, a
he the namePahudia. genus. Because Afzelia is a later homonym, adopted generic
This treatment has been either followed or ignored for a long time. In the botanical
has been treated under these three literature there are samples that one species generic names.
Fortunately, in 1935 the generic name Afzelia J.E. Sm. has been adopted and added to the Nomina Conservanda by the Botanic Congress, Amsterdam (see Sprague,
Kew Bull. 1940, 81 & 104). Hutchinson's treatment of Afzelia and Intsia (Gen. Fl.
PI. 1, 1964, 243 & 246) has been generally accepted now.
ACROCARPUS
2 547; Koord. & Val., Boomsoort. Java AcrocarpusWight ex Arnott, Mag. Zool. Bot. (1838) Bijdr.
1 (1894) 320; Polhill & Vidal in Polhill & Raven (eds.), Adv. Legume Syst. 1 (1981) 88;
Watson & Dallwitz, Gen. Leg.-Caesalp. (1983) 7. — Type species: Acrocarpus fraxinifolius
Arnott.
Trees. Stipules small, caducous. Leaves spiral, impari- or paribipinnate, pinnae and pinnules mostly oppostite; leaflets petiolulate. Inflorescence axillary, in defoliated leaf-axils, solitary, sometimes few-branched near the base and appearing fasciculate or paniculate, branches with spike-like racemes, erect, bottle-brush-like; bracts and bracteoles small, caducous. Flowers bisexual, pedicelled. Hypanthium cupular.
Calyx lobes 5, imbricate. Petals 5, imbricate. Disk cupular, completely united with the hypanthium. Stamens 5, alternating with the petals, exserted; anthers versatile,
and introrse. Ovary with a free stipe, oblong or linear, 10-20-ovuled; style stigma
the Pods elon- not sharply distinctfrom the ovary, incurved, pointed at apex. erect,
the adaxial gate, flattened, long-stipitate, narrowly winged along suture, 2-valved; valves straight, thin-coriaceous, (3-)10-18-seeded. Seeds slightly lens-shaped, smooth, not arillate, albuminous.
Distribution - So far known with one species in eastern India, Burma, Thailand,
Laos, China, and Malesia.
Note- In the original publication of the genus, Arnott (1838) distinctly cited it as
‘Acrocarpus, Wight MSS', and therefore the authorship of the genus Acrocarpus should be cited as 'Wight ex Arnott' (International Code Botanical Nomenclature,
Art. 46.3, 1988). 315 Ding Hou: Malesian Caesalpinioideae: Acrocarpus, Afzelia, Copaifera, Intsia
Acrocarpus fraxinifolius Arnott
Acrocarpus fraxinifolius Arnott, Mag. Zool. Bot. 2 (1838) 547; Wight, Icon. PL Ind. Orient. 1
(1840) t. 254; Miq., Ann. Mus. Bot. Lugd.-Bat. 3 (1867) 87; Bedd., R. Sylv. (1869) 44, t. 44;
India 2 Koord. & Kurz, For. R. Burma 1 (1877) 410; Baker in Hook. f„ R. Brit. (1878) 292; (1897) 506; Val., Bijdr. Boomsoort. Java 1 (1894) 321; Prain, J. Asiat. Soc. Bengal 66, 2
Koord., Exk. R. Java 2 (1912) 369; Koord.-Schum., Syst. Verz. 1, 1, Fam. 128 (1912) 33;
Koord., Adas Baumart. Java 1 (1913) f. 12; Heyne, Nutt. PI. Ned.-Ind. 2 (1916) 251; Craib, R.
Siam. En. 1 (1928) 507; Burkill, Diet. 1 (1935) 38; Backer & Bakh. f„ Fl. Java 1 (1964) 542;
& S.S. Larsen, Fl. Camb. Laos Vietn. 18 Brenan, Fl. Trop. E. Africa, Caesalp. (1967) 17; K. 6-9, & t. 3; 1; C.C. Hu, Fl. (1980) 76, t. 13: 7-10; Fl. Thailand 4 (1984) 50, f. 10: Reipubl.
Popul. Sin. 39 (1988) 90, t. 30; Gunn, Techn. Bull. U.S.D.A. 1755 (1991) 42, f. A-H. —
n. 845 Type: Wight 2466 (n.v.); isotype?: distributed by K under Herb. Wight (K, L), India.
291. — Mezoneurum grande Miq., R. Ned. Ind., Suppl. (1860) 108, nom. nud.; (1861) Acrocarpus
grandis (Miq.) Miq., Ann. Mus. Bot. Lugd.-Bat. 3 (1867) 87. —Lectotype (here chosen): Teijs- Sumatra. mann HB 887 (U); syntype: Teijsmann HB 862 (U), W
29 (1866) 258, nom. Cat. Acrocarpus combretiflorus Teijsm. & Binn., Nat. Tijd. Ned. Indie illeg.;
no. 908.3-1008 & Hort. Bot. (1866) 269. — Syntypes: Teijsmann s.n. (iso L, under HLB 1079),
W Sumatra.
Leaves rather large, in the herbariumoften represented by young ones or part(s) of a
and mature one; petiole and main rachis up to 80 cm long, pubescent glabrescent; basal pinnae (2 or) 3-5 pairs, secondary petiole and rachis up to c. 45 cm long, pair(s) usually shorter. Leaflets 4-7(-9) pairs per pinnule, chartaceous, ovate or acuminate base ovate-oblong, 3.5—11(—18) by 1.5—5(—8.5) cm; shortly or acute; sometimes cuneatc, obtuse, or rotund, rarely obscurely cordate, slightly asymmetric;
the midrib pubescent beneath when young, glabrescent, often slightly hairy along
when old; lateral nerves and on the petiolules, sometimesalmost glabrous 5-9(-12)
Inflorescences 32 axis per side; petiolules 2-4 mm. up to cm long, flowering pubes- cent especially towards the apical part, glabrescent; pedicels 4-10 mm, puberulous.
deflexed anthesis. 2.5-8 Flowers patent or at Hypanthium mm long. Calyx puberu- lous outside; lobes ovate or triangular, 2.5-4 mm long, glabrous inside. Petals ob-
1-2.5 both surfaces. Disk long or slightly oblanceolate, 5-9 by mm, puberulous on 15-18 anthers 2-3 hairy on the lower half inside. Stamens; filaments mm; mm long.
Pods Ovary 12-15 mm long, loosely hairy except the apical part. (8—)10—12(—17)
3-5 rather valves by (1—)1.5—2 cm (inch stipe); wing mm wide, smooth; glabrous of outside, inner surface glabrous except white hairy on and around the depressions the seeds. Seeds c. 6.5 by 5 mm. Bole Field notes - Rare, fast growing, lofty tree, 20-30(-50) by 0.6—1(—4) m.
first massive branches 10 above the columnar, often with small buttresses; (3.5-) m
lenticels. Bark whitish- ground, ultimate or young branches often with prominent
Petals dark red. Stamens with filaments grey or brown, smooth or somewhatrough.
half, the base. Seeds brown. red on the upper green at So- Distribution - See under the genus. In Malesia: Sumatra (Mt Sago, Padang,
and lok and near Batu Sangka) and Central Java (Semarang, Banjoemas Tegal).
I-L-30 & 31 and native of Cultivated in Hortus Bogoriensis sub no. XI-B-XV-11,
India.
found in Habitat - In constantly wet and fertile ground in the forest, sometimes abandoned fields, alt. 600-1200 m. 316 BLUMEA VOL. 38, No. 2, 1994
Ecology - Flowering in Feb., March, May and Dec. Fruiting in March, April,
Sept., Oct. and Nov.
Vernacular names - Sumatra: medang parrie or medang parit (Mt Sago region),
kalamparik (Padang). Java: delimas or dlimas (J).
of Notes - The authorship Acrocarpus fraxinifolius has been ascribed to Wight
(e.g. Baker, 1878), Wight & Arnott (e.g. Hutchinson, 1964) or Wight ex Arnott
(e.g. C.C. Hu, 1988). However, in 1838, when this species was published in a
Am. in Cat. 2466". It is paper by Arnott, it was cited as A. fraxinifolius, Wight n.
evident that Arnott is the sole authorof this species.
2466 listed in the of The collection of Wight no. was not lithographed copy
'Wight, Cat. Ind. PL, 1833-37' in Kew (photocopy in L), which consists of the
collection numbers 1-2403. The which bears the pages 1-144 with the specimen
Kew distribution number Wight 845 in K (one duplicate in L) may be a part of the
authentic material.
because The name Acrocarpus combretiflorus Teijsm. & Binn. is illegitimate these
two authors cited an earlier validly published legitimate name, i.e. Mezoneurum
grande Miq„ in the synonymy.
in and I have There are two syntypes of Mezoneurum grande Miq. U, chosen
collected from Suma- Teijsmann HB 887 as the lectotype. There is one specimen W
under HLB this in tra by Teijsmann (s.n. in L, no. 903.3-1086), bearing name
Miquel's handwriting, which may be a duplicate separated from one of the syntypes.
Miquel (1867) recognized the priority of the name of his new species, i.e. Me-
& Binn. and also found zoneurum grande, oxer Acrocarpus combretiflorus Teijsm.
that these two species are conspecific. He made a new combination, Acrocarpus
its grandis (Miq.) Miq. (1861), and treatedA. combretiflorus as synonym.
Baker (1878) correctly reduced Acrocarpus combretiflorus to A. fraxinifolius,
followed by Koorders & Valeton (1894), and others (e.g. Koorders, 1912). As a
result the present genus is known with only one species.
AFZELIA
Afzelia J.E. Smith, Trans. Linn. Soc. 4 (1798) 221, nom. cons., non J.E. Gmelin, 1791 (= Sey-
meria Pursh, Scrophulariaceae); Taub. in Engl. & Prantl, Nat. Pflanzenfam. 3, 3 (1894) 140,
sub Intsia; De Wit, Bull. Bot. Gard. Buitenzorg III, 17 (1941) 151; Leonard, Reinwardtia 1
(1950) 63; Brenan, Fl. Trop. E. Afr., Caesalp. (1967) 124; Cowan & Polhill in Polhill &
Raven (eds.), Adv. Legume Syst. 1 (1981) 128; Watson & Dallwitz, Gen. Leg.-Caesalp. (1983)
7. — Type species: Afzelia africana J.E. Smith ex Pers.
Pahudia Miq., Fl. Ned. Ind. 1, 1 (1855) 85; Prain, Sci. Mem. Med. Off. Army India 12 (1901) 45;
Meijer Drees, Bull. Jard. Bot. Buitenzorg III, 16 (1938)96; De Wit, Bull. Bot. Gard. Buitenzorg
III, 17 (1941) 151. — Type species: Pahudiajavanica Miq.
Trees. Stipules with their basal interpetiolar parts connate. Leaves paripinnate; leaf-
lets chartaceous, shortly petiolulate, petiolule often turned or twisted. Inflorescences
terminal often fasciculate bracts and bracteoles or axillary, racemose, or paniculate;
small, often caducous. Flowers bisexual, zygomorphic, pedicelled. Hypanthium cupular, narrowly infundibuliformor cylindric, puberulous outside, glabrous inside.
surfaces. Petals devel- Calyx lobes 4, imbricate, puberulous on both only one fully Ding Hou: Malesian Caesalpinioideae: Acrocarpus, Afzelia, Copaifera, Intsia 317
small absent. oped, large, flabellate, lower half narrowed into a claw, the others or
Disk 0. Stamens 9, usually (5—)7 fertile, almost equal, the others reduced; anthers
dorsifixed. Ovary stipitate (stipe adnate to the hypanthium), 3-8- (or more-)ovuled;
round. Pods style slender, almost as long as the stamens; stigma small, oblong,
black when obliquely oblong, or slightly rhomboid, compressed, ripe, glabrous, ovoid- 2-valved, valves thick, woody, often 3- (or more-)seeded. Seeds ellipsoid,
aril or oblong or broadly ellipsoid, smooth, exalbuminous; yellow, orange, red, half of its often 2-lobed, covering the seed for up to or more length.
Distribution- About 12 species, in tropical Africa, Asia and S China; two species
in Malesia.
Habitat - In forest at low and medium altitudes, 0-400(-1400) m.
underthe It is diffi- - allied to Intsia note latter). Note Afzelia is very closely (see
cult to identify incomplete or sterile herbarium collections of these two genera.
KEY TO THE MALESIAN SPECIES
la. Leaves usually pubescent, sometimes densely pubescent on the petiole, rachis, leaflets. petiolules, and especially on the midrib of the lower surface of Petals
not papillate. Stamens with filaments united along the lower (1.5—)2—2.5 cm 1. A. javanica
b. Leaves often glabrous, sometimes sparsely (very rarely densely) pubescent on
midrib of the lower the petiole, rachis, petiolules, and especially on the surface
the inner surface. Stamens with of leaflets. Petals often papillate at least on
filaments united at the base (or almost free) 2. A. rhomboidea
1. Afzelia javanica (Miq.) Léonard
Afzelia javanica (Miq.) Ldonard, Reinwardtia 1 (1950) 63; Gunn, Techn. Bull. U.S.D.A. 1755
Koord. & (1991) 252, f. G. — Pahudia javanica Miq., Fl. Ned. Ind. 1, 1 (1855) 86; Val., Bijdr.
Boomsoort. Java 2 (1895) 28; Prain, Sci. Mem. Med. Off. Army India 12 (1901) 46, f. 6-11;
Bot. Heyne, Nutt. PI. Ned.-Ind. 2 (1916) 239; ed. 2, 2 (1927) 736; Meijer Drees, Bull. Jard.
Buitenzorg III, 16 (1938) 98, f. 2a; De Wit, Bull. Bot. Gard. Buitenzorg III, 17 (1941) 145. —
Pahudia javanica subsp. javanica; Backer & Bakh.f., Fl. Java 1 (1964) 530. — Pahudiajavanica
subsp. eujavanica De Wit, Bull. Bot. Gard. Buitenzorg III, 17 (1941) 146, f. 2: 1. — Afzelia
javanica subsp. javanica; L6onard, Reinwardtia 1 (1950) 63. —Type: Th. Horsfield 'L 147' (K,
iso BM, photo of both types in L), Central Java, near Prowoto.
290. — Pahudia Intsia puberula Miq.,Fl. Ned. Ind. Suppl. (1860) 107, nom. nud.; (1861) puberula
(Miq.) Meijer Drees, Bull. Jard. Bot. Buitenzorg III, 16 (1938) 97; De Wit, Bull. Bot. Gard. Sumatra, Buitenzorg III, 17 (1941) 145. —Type: Diepenhorst s.n. (U, sh. 038678, photo in L),
Priaman.
Pahudiajavanica subsp. longiflora De Wit, Bull. Bot. Gard. Buitenzorg III, 17 (1941) 146, f. 2: 2.
— — bb Afzelia javanica subsp. longiflora (De Wit) Leonard, Reinwardtia 1 (1950) 63. Type:
31600(iso L), Sumatra, East Coast.
also Young twigs, rachides, leaflets especially on the midrib of the lower, sometimes
sometimes or glabrescent. Leaves on the upper,surface, sparsely, densely pubescent
5- and rachis Leaflets (4-) or 6(-8)-jugate; petiole up to c. 20(—30) cm. oblong-
elliptic or -ovate, rarely ovate or broadly elliptic, 6—9(—16) by 2.5-4(-5.5) cm; apex 318 BLUMEA VOL. 38, No. 2, 1994
shortly acuminate, obtuse, rarely retuse; base rounded or obtuse, slightly subcordate,
7-14 veins 2-3 mm. or acute; lateral nerves per side; loosely reticulate; petiolules
Inflorescences with main axis often 4—12(—24) cm, pubescent; bracts and bracteoles
8-12 5-7 3-5 mm long; pedicels mm, puberulous. Hypanthium mm long. Calyx 8-10 lobes broadly elliptic or suborbiculate, 6-10 by 4-7(-9) mm. Developed petal
Stamens: filaments united the lower by 7-9 mm. (4.5—>5.5—6.5 cm long, along
towards the base inside anthers 1- (1.5—)2—2.5 cm, glabrous except thinly hirsute;
6 1.5 mm long; staminodes c. 4 mm, glabrous. Ovary c. mm long, densely puberu-
5-7.5 1.5-2 thick. lous along the margin; style 4-5.5 cm. Pods 7-14 by cm, cm
thick. Seeds 1-8, 2-3 by 1.5-2.2 cm, c. 1.3 cm
- m d.b.h., often much smaller. But- Field notes Tree up to 42 m tall, 0.7(—1.3)
8 cm thick. Bark thin, with brown or tresses up to 5 m high, 1.25 m wide, and grey
bark 10-15 mm thick, white or brown. Petals white. black, without grooves; inner
red. Aril yellow, orange, or
Distribution- Sumatra: Atjeh and E Coast. W & C Java: Preanger, Panandjung,
Prowoto (= Prawata), Banjoemas, Cheribon, and Pekalongan.
Habitat - In primary and secondary forests, in dry places, rarely occurring on
alt. limestone, usually found at 0-100 m, sometimes up to 400(-800) m
Ecology - Flowering in April, Oct. Fruiting in Jan., Feb., March, May, Aug.
- ketaroem kataroem Vernacular names Sumatra: hataroem or (Batak), (M), mer- bau (Atjeh). Java: djoelang perit (J), kidjoelang or djoelang (S & J), kidjoelang tau- doek or k. kapas (S).
Note - The developed petal often falls off at anthesis, so it is often lacking in the herbarium.
2. Afzelia rhomboidea (Blanco) Vidal
Afzelia rhomboidea (Blanco) Vidal, Cat. PI. Prov. Manila (1880)28; Phan. Cuming. Philipp. (1884)
26, 110; Rev. PL Vase. Filip. (1886) 117; Leonard, Reinwardtia 1 (1950) 63; Gunn, Techn.
Bull. U.S.D. A. 1755 (1991) 256, f. B. — Eperua rhomboideaBlanco, Fl. Filip., ed. 2 (1845)
192. — 260; ed. 3, 2 (1878) 119, t. 281. — Intsia rhomboidea Kuntze, Rev. Gen. PI. 1 (1891)
Pahudia rhomboidea Prain, Sci. Mem. Med. Off. Army India 12 (1901) 46; Merr., Philipp. J.
Sc. 1 (1906) Suppl. 63; 5 (1910) Bot. 41; Whitford, Bull. Bur. For. Philipp. 10 (1911) 39;
Men., Fl. Manila (1912) 236; Sp. Blanc. (1918) 171; Enum. Philipp. Fl. PI. 2 (1923) 257;
Univ. Calif. Publ. Bot. 15 (1929) 101 (as Afzelia javanica Miq.); Meijer Drees, Bull. Jard. Bot.
Buitenzorg III, 16 (1938) 99; De Wit, Bull. Bot. Gard. Buitenzorg III, 17 (1941) 147. —
Neotype (here chosen): Merrill Sp. Blanc, no. 862 (L; iso BO), Luzon, Laguna Prov. 1 369; Merr., Blanc. (1918) 171. Eperuafalcata auct. non Aublet: Blanco, Fl. Filip. ed. (1837) Sp.
Intsia acuminata Men., Publ. Gov. Lab. Philipp. 17 (1904) 20; Enum. Philipp. Fl. PI. 2 (1923)
257. —Type: Merrill 1108 (iso K), Luzon.
Afzelia borneensis Harms in Fedde, Rep. Sp. Nov. 14 (April 1916) 256; 15 (1917) 19; Leonard,
Reinwardtia 1 (1950) 63; Meijer, Field Guide Trees W. Mai. (1974) 179, f. 41; Cockb., Trees
J. 11 Bot. 86. Sabah 1 (1976) 157, f. 32. — Pahudia acuminata Men., Philipp. Sc. (June 1916)
— Afzelia acuminata (Men.) Harms in Fedde, Rep. Sp. Nov. 15 (1917) 19. — Pahudia bor-
neensis Men., J. Sir. Br. R. As. Soc. 76 (1917) 84; Bibl. Enum. Born. PL (1921) 297; Meijer
Gard. Drees, Bull. Jard. Bot. Buitenzorg III, 16 (1938) 100, f. 2c; De Wit, Bull. Bot. Buitenzorg
III, 17 (1941) 148, f. 2: 4. — Lectotype (vide Harms, 1917): Hose 93 (iso L), Sarawak, Baram. DingHou: Malesian Caesalpinioideae: Acrocarpus, Afzelia, Copaifera, Intsia 319
Pahudia rhomboidea var. praetermissa De Wit, Bull. Bot. Gard. Buitenzorg III, 17 (1941) 151, f. 2:
Prain 1 63. — 3a. — Afzelia rhomboidea var. praetermissa Leonard, Reinwardtia (1950) Syn-
bb East types: Clemens 21289 (L), Sarawak, Kapit; 8981 (BO, n.v.), Sumatra, Coast; Exp.
Nieuwenhuis 1424 & 1509 (BO, n.v.), Kalimantan, Bloe-oe; Teijsmann 8249 (BO, n.v.), Kali-
mantan, Mt Singkadjang.
Young twigs, rachides, petioles, petiolules, and leaflets often glabrous, sometimes
the midrib of the lower sparsely, very rarely densely pubescent, especially on
the surface. Leaves 4- and rachis (sometimes also on upper) (3-) or 5-jugate; petiole
Leaflets 10—14(—19) cm. ovate, elliptic, 3.5—10(—15) by 2.5-4.5(—6) cm; apex
(shortly) acuminate; base acute or obtuse, rarely more or less truncate or shallowly
concave; lateral nerves 5-8 per side and veins rather closely reticulate; petiolules
both 1.5-4.5 mm. Inflorescences with main axis up to 12—14(—15) cm long, main axis and branches puberulous; pedicels 4.5-5.5 mm. Hypanthium c. 3 mm long.
5-9 7-10 Calyx lobes broadly ovate or subrotund, 5-12 by mm. Developed petal
filaments 4.5-6 united the base almost by 5-9 mm. Stamens: cm, at (or free),
slightly hairy near the base; anthers c. 1.5 mm long; staminodes (0.5—) 1—3 cm,
sparsely hairy. Ovary 1.5-4 mm long, puberulous along the margin; style two
the base. Pods kinds: long (40-50 mm), or very short (1-2 mm), slightly hairy at
thick. Seeds 1.7-3 1.5-2.5 1 9—12(— 20) by 5 —7(— 10) cm, c. 1.5 cm by cm, c. cm
thick.
Field notes - Tree 25-30(-36) m tall, 0.4-0.8(-1.2) m d.b.h. Bole cylindric.
Buttresses sometimes present up to 2 m high, 1-2.5 m wide and c. 8 cm thick. Leaves
2 fall once a year, but not at the same time in all individual trees. Flowering during or
3 weeks, probably annually, not after falling of leaves. Petals yellowish red, red or
(older) dark red. Aril orange or yellow, orange-red.
Distribution - Malesia: Sumatra (East Coast and Palembang), Sabah (Kalabakan,
Mt Kinabalu, Mostyn, and Tawau), Kalimantan (Berouw: Mt Has Bungaan), and
Philippines (N Luzon to Mindanao).
Habitat- In primary forests, occasionally occurring on limestone or sandstone,
350 found at 1400 Kinabalu: SAN up to m alt., once c. m (Mt 42722, L).
Ecology - Flowering and fruiting almost all year round.
Uses - Valuable timbertree.
Vernacularnames - Sumatra: merbau tandoek (Palembang). Sabah: merbau tan-
under Pahudia doek (Tawau). Philippines: see Merrill (1923: 257, rhomboidea).
- borneensis Harms 1916: citedwith Notes Afzelia was published by (April 256), two specimens: Hose 93 (in fl„ L) and H. Winkler 3236 (in buds). One year later
based Harms (1917: 19) stated that his species was on Hose 93, so this specimen should be treated as the lectotype (vide Intern. Code Bot. Nom., Art. 7.4 & 7.5,
1988).
Pahudia acuminatawas described by Merrill (1916: 86) based also on Hose 93.
1916 1917: It was published in June (not March) (vide Merrill, 84).
Harms (1917: 15) found that the above two species were based on the same col-
of Merrill's lection, i.e. Hose 93, but misquoted the publication date species as one combination, i.e. monthearlier than his, so he made an unnecessary Afzelia acumi- nata (Merr.) Harms. 320 BLUMEA VOL. 38, No. 2, 1994
had been treated distinct Afzelia borneensis and A. rhomboidea as two species by
using the length of styles as the main character: style 1-2 mm in the former against
40(-50) mm in the latter. However, after having examined many collections of these
two species, their flowers appeared to be heterostylous. These two species are con-
specific.
COPAIFERA
Copaifera L., Sp. PI. ed. 2 (1762) 557, nom. cons.; De Wit, Webbia 9 (1954) 459; Leonard, M6m.
Acad. R. Belg. CI. Sci. 30 (1957) 84; Hutch., Gen. Fl. PI. 1 (1964) 253; Cowan & Polhill in
Polhill & Raven (eds.), Adv. Legume Syst. 1 (1981) 132; Watson & Dallwitz, Gen. Leg.-
Caesalp. (1983) 20. — Type species: Copaifera officinalis (Jacq.) L.
Pseudosindora Symington, Proc. Linn. Soc. Lond. 155 (1944)285; Leonard, M6m. Acad. R. Belg.
CI. Sci. 30 (1957) 86; Cowan & Polhill in Polhill & Raven (eds.), Adv. Legume Syst. 1 (1981)
132. — Type species: Pseudosindorapalustris Symington.
Trees. Stipules caducous. Leaves imparipinnate, petiolate; leaflets alternate, petiolu-
late. Inflorescences axillary, paniculate; bracts and bracteoles small, caducous. Flow-
0. lobes im- ers bisexual, actinomorphic, pedicelled. Hypanthium Calyx 4, narrowly
bricate or subvalvate. Petals 0. Disk 0. Stamens 10, free; anthers dorsifixed. Ovary
stipitate, 2-ovuled; style slender; stigma small, capitellate. Pods ellipsoid, rather
smooth, 2-valved, valves thick coriaceous, 1- or 2-seeded. Seeds oblong, arillate,
exalbuminous.
Distribution - Species 25-30, mostly in tropical America, 4 in tropical Africa,
and 1 in Malesia (Borneo).
Habitat- See under the species.
Notes - The genus Copaifera is allied to Sindora but differs from it by the alter-
nate leaflets (against opposite leaflets) and lacking petals (against with a well devel-
oped petal). It is also related to Crudia but differs from it by the paniculate inflores-
cences, the lack of a turbinate hypanthium, and distinctly arillate seeds.
De Wit (1954: 459-464) compared Pseudosindora with the related genera Co- paifera, Crudia, and Sindora. He reducedPseudosindora to Copaifera.
Leonard (1957: 84) expressed the opinion that it is preferable to maintain Pseudo- sindora, because it differs from Copaifera by various characters, especially by the
ranks. have examined arrangementof flowers in more than two I flowering speci- mens of Copaifera ; the flowers are not always distichously arranged but are some- times shown in more than two ranks.
Leonard (1957: 84 & 86) studied also the seedlings offour species: one American
of the with the first leaves species, Copaifera officinalis L., type species genus, two
three African with the first leaves The fruits alternate; ones two opposite. open com- pletely in two valves in the American species but dehisce from the apex towards 1/2-
3/4 of their length in the African ones.
of The seedlings ofthe Malesian species, Copaifera palustris, are similar to those
American the African ones, while the dehiscence of the fruits is like that of the one
(cf. Symington, 1944: 285-288, f. 1 & 2).
Pseudosindora in the of Hutchinson (1964: 253) also placed the genus synonymy
Copaifera. Ding Hou: Malesian Caesalpinioideae: Acrocarpus, Afzelia, Copaifera, Intsia 321
Cowan and Polhill (1981: 132) kept Pseudosindora and Copaifera as two distinct
former distinct from in generabut stated that the is: "Debateably Copaifera, differing
the spirally arranged flowers and larger fruits with medially inserted seeds." I have examined some flowering collections (e.g. Beccari PB 3915, L, cited also by Sy-
mington; S 12268, L), which bear the young flower buds more or less distichously
arranged; however, the old buds appear more or less spirally arranged due to the
slightly twisted rachides. The infructescences are slightly twisted so the fruits or their
which scars appear spirally arranged. The ovary often contains 2 ovules are medially
the seeds less inserted. A fruit may contain 1 or 2 developed seed(s); are more or
in the half when medially inserted especially when young, but are upper or subapical old.
of flowers and seeds features but alter The arrangements the are developmental
with with De Wit and Hutchinson in the Pseudosindora age. I agree reducing genus
to Copaifera.
Copaifera palustris (Symington) De Wit
Copaifera palustris (Symington) De Wit, Webbia 9 (1954) 462. — Pseudosindorapalustris Syming-
& 2. — C. O. Flemmich FMS 32600 ton, Proc. Linn. Soc. Lond. 155 (1944) 285, f. 1 Type:
(KEP, n.v.), Brunei.
Stipules auriculate to subfalcate, up to 1.5 by 1 cm. Leaves (2-)4(-6)-foliolate; peti-
often ole and rachis 4.5-12 cm, glabrous. Leaflets coriaceous, elliptic-oblong, un-
base equally sided, 5-9(-14) by 3.5-6(-8) cm; apex acute or shortly acuminate;
sometimes the rotund; glossy, pellucid-dotted, glabrous except sparsely hairy on
midrib the base the lower distinct visi- or at on surface, glabrescent; nerves many, or ble on both surfaces; often with 1-3 elliptic, slightly depressed glands towards the basal part of the margin; petiolule 4-7 mm. Inflorescences 4-11 cm long, pubescent; pedicels 3-4 mm, puberulous. Calyx lobes elliptic, 4-7.5 by 2-4 mm, puberulous outside, densely tomentose inside. Stamens alternately long and short; filaments7-
2-3 10 mm, glabrous; anthers c. 2 mm long. Ovary by 1-1.5 mm, pilose along the
and 1-2 4.5-5 Pods 4.5-7.5 ventral suture on the stipe; stipe mm; style mm. by
brown aril 2- 3.5-4 cm. Seeds 2.4-2.7 by c. 1.5 cm, glossy or black; narrowly lobed, enclosing the seed.
Field notes - Tall unbuttressed tree, up to 30(-39) m high, up to 40(-180) cm d.b.h.; bark minutely fissured. Flowers pearl-fawn coloured. Aril pink.
Distribution - Malesia: Brunei; Sabah (Temburong); Sarawak (Igan near Sibu,
Betong, Miri, Binatong, and Baram); Kalimantan (Sekadau).
- forest the found in Habitat In freshwater peat swamp near coast, sometimes also dry forest, once found in secondary heath forest, 0-30 m alt.
Ecology - Flowering in April, Oct. Fruiting in July.
Vernacular names - Emputir (Dajak-Malay), sepetir (Brunei & M), sepetir paya
(Baram & Miri), tepih (Miri).
Uses - Symington (1944: 288) reported that the "timberis neither durablenor dec-
is for the of boats." Because ofits moderate orative and the only recorded use masts hardness, good shape, and local abundance, he suggested that it may be used in ply- wood manufacture. It is also used for roof tiles. 322 BLUMEA VOL. 38, No. 2, 1994
INTSIA
Intsia Thouars, Gen. Nov. Madag. (1806) 22; Prain, Sci. Mem. Med. Off. Army India 12 (1901)
43; Meijer Drees, Bull. Jard. Bot. Buitenzorg III, 16 (1938) 87; De Wit, Bull. Bot. Gard. Buiten-
17 Reinwardtia 1 Gen. Fl. PI. 1 zorg III, (1941) 139; L6onard, (1950) 61; Hutch., (1964)
245; Cowan & Polhill in Polhiil & Raven (eds.), Adv. Legume Syst. 1 (1981) 128; Watson &
Dallwitz, Gen. Leg.-Caesalp. (1983) 35. — Intsia sect. Euintsia Taub. in Engl. & Prantl, Nat.
Pflanzenfam. 3, 3 (1894) 140, nom. illeg. — Afzelia auct. non J.E. Smith: Koord. & Valeton,
Bijdr. Boomsoort. Java 2 (1895) 30; Prain, J. Asiat. Soc. Bengal66, 2 (1897) 207. — Type
Thouars DC. Intsia species: Intsia madagascariensis ex (vide Hutch., 1964) = bijuga (Colebr.)
Kuntze.
Ann. Bot. 4 1789. Macrolobium Zipp., nom. nud., pro syn., Walp., Syst. (1857) 610, non Schreb.,
Trees often with buttresses. Stipules intrapetiolar connate. Leaves paripinnate. Leaf-
lets chartaceous to subcoriaceous, usually with 1 or 2 small crateriform glands at the
basal part on lower surface; petiolules twisted. Inflorescences terminal or axillary,
bracts caducous. Flowers simply racemose, often fasciculate, or paniculate; bisexual,
zygomorphic. Hypanthium cupular, narrowly infundibuliform or cylindric. Calyx
lobes 4. Petals: only 1 fully developed, flabellate, lower half narrowed into a claw,
the others rudimentary or absent. Disk 0. Stamens 3 or very rarely 4 fertile; stami-
nodes 4-7; filaments and staminodes connate at the base; anthers dorsifixed. Ovary
stipitate (stipe adnate to the hypanthium except the apical part), puberulous; style
slender; stigma small, capitellate. Pods oblong, rarely obcordate, straight, or falcate,
flattened, glabrous, dehiscent, 2-valved (valves leathery or slightly woody), often 3-
(or more-)seeded. Seeds ovoid, oblong, discoid, or sublenticular, flattened, not aril-
late, scurfy, exalbuminous.
Distribution - Two or more species, from Madagascar, the islands of the Indian
Ocean, tropical Asia, through Malesia to N Australia, Melanesia and Micronesia.
In Malesia two species occur, see Figure 1 (cf. De Wit, 1941: f. 1; Whitmore, 1972: f. 9).
Habitat - The of Intsia of the Malesian species are 'long-lived secondary' type
(see below), occurring from coastal, salty regions up to about 1000 m altitude.
Regeneration - Seeds have a period of dormancy, requiring pre-germination treat- ment (e. g. puncturing of the testa) in silviculture. Seedfallmight be very good, but abundant regeneration takes place only in forest gaps, after fires, or after felling.
Saplings tend to loose branches and fail to develop a leading shoot. Growth is slow.
Because of lack of regeneration under closed canopy, the distributionover diameter classes is not regular, but reflects earlier disturbances in the forest. Zieck (internal report West New Guinea) described gregarious occurrences of Intsia bijuga on steep slopes of raised coral reef almost without any soil, but this species also occurs in swampy, even salty, habitats.
Intsia reaches because it has palembanica generally larger sizes, maybe a narrower
and soils. Vanimo ecological amplitude, avoiding wet very dry In (Papua New Guinea) good stands have been produced after forest fires-apparently after a good seed-fall- in the 1880's, with 23% net volume of the total stand. Rotationwould thus at least be 80 years.
Internationaltrade name - Merbau (in Papua New Guinea: Kwila). Ding Hou: Malesian Caesalpinioideae: Acrocarpus, Afzelia, Copaifera, Inlsia 323
Kuntze and Intsia Fig. 1. Distribution range of Intsia bijuga (Colebr.) (solid line) palembanica Miq.
(dashed line).
Uses - Trees of exploitable size (diam. 60 cm and up) have rather little, white to
yellowish brown sapwood, which is sensitive to early deterioration, e.g. by powder-
beetles and The heartwood is reddish dark post (Lyctis ) fungi. brown, weathering to
brown, heavy (S.G. 0.71-1.05, airdry), hard, very strong, durable, resistent to ter-
mites but not to marine borers. Yellowish deposits in the vessels are often con-
spicuous, often showing on longitudinal surfaces, water-soluble and therefore cause
staining of non-sealed timbersurfaces exposed to water. Silica absent, but saw teeth
can become gummy. Shrinkage and movement very low. Grain straight or slightly
interlocked. The wood finishes well. Pre-boring for nailing and screwing is advis-
able.
of both indoors and outdoors not in Intsia timber has a wide range uses (if contact
with seawater), e.g. (heavy) constructions, flooring, doors, posts, poles, sleepers,
etc.
Literature - Den Berger, Meded. ProefstationThee 97 (1926) 53; Glifford, Tim- ber identification (1957) 122; Eddowes, Commercial timbers of PNG (not dated) 20,
commercial timbers of PNG 116; Species notes on the major (not dated, not paged);
Foxworthy, Malayan Forest Rec. 3 (1927) 88; Johns, Blumea 31 (1986) 342; Lane-
Poole, Forest resources Papua & New Guinea (1925) 92b; Versteegh, Meded. LH
Wageningen 17-19 (1971)) 40; Whitmore, Tree Fl. Malaya 1 (1972) 262; Trop. rainforests Far East (1975) 182. -— W. Vink.
- is allied to It differs from the latter Notes The genus Intsia very closely Afzelia.
mainly by having 3 or very rarely 4 (against usually 7) fertile stamens and not arillate
(against prominently arillate) seeds.
There are many Malesiancollections of the two widely distributed, very variable
species treated here. Sterile material is very difficult to identify. 324 BLUMEA VOL. 38, No. 2, 1994
KEY TO THE SPECIES
la. Leaves 2- (or 3-)jugate, rarely 1-jugate towards the top of twigs or below the in-
florescences. Hypanthium 6—10(—16) mm long, usually about the same length
as the calyx lobes. Style c. 4 cm. Pods 8.5—15(—28) by 4—5(—7.2) cm
1.1, bijuga
b. Leaves 4-jugate, rarely associated with some 2-, 3- or 5-jugate ones. Hypanthium
than the lobes 6-8 2- 3-4 mm long, shorter calyx (usually mm long). Style
3.5 cm. Pods 15—19(— 40) by 6-7(-9) cm 2. I. palembanica
1. Intsia bijuga (Colebr.) Kuntze
Intsia bijuga (Colebr.) Kuntze, Revis. Gen. PI. 1 (1891) 192; Merr., Int. Rumph. (1917) 255; Sp.
Blanc. (1918) 171; Brown, Minor Prod. Philipp. For. 2 (1921) 393, f. 1; Merr., En. Philipp.
Fl. PI. 2 (1923) 257; Kanehira, Bot. Mag. Tokyo 45 (1931) 285; Fl. Micron. (1933) 142, f. 48;
Burkill, Diet. 2 (1935) 1264; Meijer Drees, Bull. Jard. Bot. Buitenzorg III, 16 (1938)89, f. lb,
incl. f. hirsuta Meijer Drees and f. glabraMeijer Drees; De Wit, Bull. Bot. Gard. Buitenzorg III,
17 (1941) 140, f. 1; Holthuis & Lam, Blumea 5 (1942) 192, as f. glabra Meijer Drees; Kane-
hira & Hatusima, Bot. Mag. Tokyo 56 (1942) 362; Corner, Ways. Trees ed. 2, 1 (1952) 396;
Walker, Important Trees Ryukyu Is. (1954) 124, f. 65; Backer & Bakh. f., Fl. Java 1 (1964)529;
Whitm., Guide For. Brit. Solom. Is. (1966) 80; Brenan, Fl. Trop. E. Afr., Caesalp. (1967) 128,
f. 23; Zuijderh., Blumea 15 (1967) 425; Hatusima, Fl. Ryukyus (1971) 343; Whitm., Tree Fl.
Mai. 1 (1972) 262, f. 9; Cockb., Trees Sabah 1 (1976) 170, f. 36; Walker, Fl. Okinawa S.
Ryukyu Is. (1976) 542, f. 72; Verde., Man. N. G. Legumes (1979) 91; K. & S.S. Larsen, Fl.
Camb. Laos Vietn. 18 (1980) 144, t. 25: 5-9; Fl. Thailand 4 (1984) 125, f. 31: 5-9, pi. 4: 4;
A.C. Sm„ Fl. Vitien. Nova 3 (1985) 134, f. 26; Gunn, Techn. Bull. U.S.D.A. 1755 (1991)
258, f. B, C, E-H. — Macrolobium bijugum Colebr., Trans. Linn. Soc. 12 (1818) 359, t. 17.
— A. U.S. — Outea bijuga DC., Prod. 2 (1825) 511. Afzelia bijuga (Colebr.) Gray, Expl.
Exped., Phan. 1 (1854) 467, nom. illeg., non Afzelia bijuga (Willd.) Spreng., 1827 (= Erio-
glossum cauliflorum Guill. & Perr., Sapindaceae); U.S. Expl. Exped., Atlas Phan. 1 (1856)
t. 51; Seemann, Fl. Vitiensis (1865) 69; Kurz, For. Fl. Burma 1 (1877) 412; Baker in Hook, f.,
Fl. Brit. Ind. 2 (1878) 274; Kuntze, Revis. Gen. PI. 1 (1891) 192; Koord. & Val„ Bijdr. Boom-
soort. Java 2 (1895) 31; Prain, J. Asiat. Soc. Bengal 66, 2 (1897) 208; Pulle, Nova Guinea 8
(1912) 650; Ridley, Fl. Mai. Pen. 1 (1922) 639. — Type: No collection cited; based on a plant
cultivated in the Calcutta Botanic Garden, no certainly authentic material seen; based on Cole-
brooke's descr. & pi. (see under Note).
Intsia madagascariensis Thouars ex DC., Prod. 2 (1825) 509. — Type: Based on descr. of Intsia
Thouars (1806).
Intsia? amboinensis DC., Prod. 2 (1825) 509; Miq., Fl. Ned. Ind. 1, 1 (1855) 80; Fl. Ned. Ind.,
Suppl. (1860) 107 & 288. — Metrosideros amboinensis Rumph., Herb. Amb. 3 (1750) 21,
t. 10. — Macrolobium amboinense Teijsm. ex Hassk., Abh. Naturf. Ges. Halle 9 (1866) 189.
— Metrosiderosamboinensis Type: Based on descr. and pi. of Rumph.
Jonesia triandra Roxb., Fl. Ind. ed. 2, 2 (1832) 220. — Saraca triandra Baker in Hook, f., Fl. Brit.
cultivated in the Calcutta Ind. 2 (1879) 272. — Type: Plant from the Malay Archipelago, maybe
Bot. Gard.; authentic specimen(s) not seen; based on descr.
Eperua decandra Blanco, Fl. Filip. (1837) 368; ed. 2 (1845) 259; ed. 3, 2 (1878) 118. — Neotype
(here chosen): Merrill Sp. Blanc, no. 376 (L; iso in BO), Malampaya Bay, Palawan.
Jonesia monopetala Hassk., Retzia 1 (1855) 199. — Pahudia hasskarliana Miq., Fl. Ned. Ind. 1,1 (BO, n.v.), Java: Bantam. (1858) 1080, nom. illeg. —Type: A. Boachi s.n.
— 'L 187' Saraca obtusifolia Miq., Fl. Ned. Ind. 1, 1 (1855) 85 & (1858) 1080. Type: Horsfield
(K, photo in L; iso U, photo in L), Java (Banjoemas). Ding Hou: Malesian Caesalpinioideae: Acrocarpus, Afzelia, Copaifera, Intsia 325
Intsia moelebei Vieillard, Bull. Soc. Linn. Normand. 9 (1865) 339. — Type: Vieillard 386 (iso in
L), New Caledonia.
12 Intsia retusa (Kurz) Kuntze, Revis. Gen. PI. 1 (1891) 192; Prain, Sci. Mem. Med. Off. Army India
(1901) 44; Merr., Philipp. J. Sc. 11 (1916) Bot. 85; Bibl. Enum. Born. PI. (1921)296; Philipp. J.
Sc. 30 (1926) 399; Burkill, Diet. 2 (1935) 1265; Meijer Drees, Bull. Jard. Bot. Buitenzorg III,
16 (1938) 88, f. la. —Afzelia retusa Kurz, J. Asiat. Soc. Bengal 42, 2 (1873) 73; Fl. Burma 1
(1877) 412; Baker in Hook.f., Fl. Brit. Ind. 2 (1878) 274; Prain, J. Asiat. Soc. Bengal 66, 2
(1897) 207 & 494; Ridley, Fl. Mai. Pen. 1 (1922) 639. — Type: From the Andamans, maybe
collected by Kurz, no specimen cited; authentic material not seen, based on description.
Intsia tashiroi Hayata, Ic. PI. Formos. 3 (1913) 85. — Type: Y. Tashiro s.n. (n.v.), S Ryukyu:
Yaeyama Archipelago.
Intsia bijuga f. hirsuta Meijer Drees, Bull. Jard. Bot. Buitenzorg III, 18 (1938) 91. — Syntypes: bb
2990 (BO, n.v.), Celebes: Saleijer; bb 3918 (L), Celebes: Moena; bb 3712 (BO n.v.),
Moluccas: Temate.
towards the of below the inflores- Leaves 2- (or 3-)jugate, rarely 1-jugate top twigs or
and rhachis Leaflets cences; petiole (1.5—)2.5—11.5 cm long, glabrous. (broadly)
elliptic, ovate, obovate, rarely suborbiculate, (2.5—)5—10(—18) by (1.5—)4—6(—12)
rounded, sometimes notched; cm; apex obtuse, shortly acuminate, rarely slightly 2-6 Inflo- base acute, cuneate, or obtuse, often slightly asymmetric; petiolules mm.
less rescences up to 10(—17) cm long, pubescent, glabrescent, or more or glabrous;
about the the lobes. pedicels 7-12 mm. Hypanthium usually same length as calyx
Petal: lamina Calyx lobes elliptic, 6—10(—16) by 4-5(-7.5) mm. 6-10(-30) by 10-
5 Fertile filaments 3 anthers 2 15(—35) mm; claw c. mm long. stamens: c. cm; c. mm
10 7.5 4 Pods long; staminodesup to mm. Ovary c. mm long; style c. cm. 8.5—15(—
28) by 4-5(-7.2) cm. Seeds 2-3.5 cm long and wide, c. 0.8 cm thick.
- Field notes Tree up to 40 m high; bole straight, sometimes crooked, up to 24 m
long and 1 m d.b.h., but usually smaller. Buttresses when present up to 2(-4) m
with small shallow circular high, extending outwards c. 2 m. Bark covered depres-
Fertile with red sions. Petals white, later turning pink, red, or purple. stamens or
Pods brown black when purple filaments. Style red or purple. or ripe.
Distribution - As under the genus.
Cultivatedin Africa (Zanzibar and Tanzania, fide Brenan, 1967), in India (Calcut-
ta Bot. Gard.), and in Java (Hort. Bog. sub no. I-I-2, died in 1959; I-C-16a).
Habitat - Along (sandy) sea coasts, edges of rivers, tidal or temporarily inundated
places with (salty) water, back-mangroves, in primary and secondary forests, in Kali-
recorded from New Guinea rather mantan once foundon acid sandy soil; western as
BW from common (Adi I., BW 9825, L) or very common (Japen I., 7009, L), sea-
level up to c. 600 m alt.
Ecology - Flowering and fruiting all year round.
- Uses See under the genus.
- merkau Pe- Vernacular names Sumatra: merkau(M), ajer (East Coast). Malay
ninsula: ipil (or ipeh), ipil tandok, malapari, merbau, merbauayer, merbauchangkat,
merbau laut (M). Java: merbaoe, merbau, merbo, taritih (Sundanese). Lesser Sunda
Brunei: Sabah: merbau Islands: ibla (Sumba), ipi (Flores). ipil-ipil (M). epil, paya,
ipil (Bajan), moebau (M), talolonandim(Beaufort). Kalimantan: ipil merbau suglai
1923: Celebes: (Sesajap). Philippines (see Merrill, 257). bajani (bajang or bajam)
(Makassar), gefi (or ogifi), ipi, ipilo (Moena), ipi (Koeli & Taa), kintom (Banggai), 326 BLUMEA VOL. 38, No. 2, 1994
Ian ggiri (Tobela-To Tambei), loroeroe (Tobela-To Padoe). Moluccas: dowora(Mo-
rotai & fraa besi Ternate-Tidore), (Kai), kajoe merah or kajoe besi panti (Ceram),
sikata'a (Talaud), kajoe tea (or k. besi) (Soela), aakonal, tahai hoboi or takai koboi
(= aakomal = besi koening) (Aru). New Guinea: arir (Wanigela), ariri (Onjob), babibi
(Sentani), babrie (Sentani), bat (Kemtoek), bauw (Atam), bendoro (Oro Bay &
bon duhum ferraai Orokaiva), (Madang), (Mawan), epna (Labu), (Kowiai), ganam
(Jal), gommagome(Kawa), haboe (Noemfoor), haero (Goaribari, Janbwank, Kai-
gorin), jep (Bembi), kaboei (Biak), kwila (Pidgin), amele, mep, milimbu (Faita),
paseh (Asmat), patoem (Njou), pian (Amberbaken), pota (Mimika), rang & raung
(Sko), rong (Ambai), seka (Manikiong), tangibe (Sko), wohne (Waskuk), yambwan
(Dunpu). New Britain: bana (Garumaia).
Notes - Colebrooke (1818) published his new species Macrolobium bijugum,
based on a plant of unknown origin and date of introduction, cultivated in the Cal-
cutta Botanic Garden. He gave a detailed description and one well-prepared plate
of and of the but did cite (with drawings habit, flowers, pod, seed) plant, not any
collection. I have not seen any publication indicating that the authentic material has
ever been preserved or is extant. However, one well-preserved specimen, i.e. Wallich
Cat. no. 5823A (K-W; K, photo in L; on IDC microfiche) is from the Calcutta Bot.
Gard. (cf. Brenan, 1967: 130). From his description and plate the identity of this
species had been correctly interpreted by Kuntze (1891), followed by others, as be-
longing to Intsia, as I. bijuga (Colebr.) Kuntze.
Saraca obtusifolia Miq. was treated as a synonym ofSaraca declinata (Jack) Miq.
1967: The of this in by Zuijderhoudt (Blumea 15, 419). type species is Kew bearing
threelabels: Javanicum Horsfield 1) 'Herb. Dr. L 187'; 2) name of this new species in Miquel's handwriting; and 3) a special printed label of Kew for the specimens col- lected by Thomas Horsfield from Java (in 1802-1818). This specimen had been an-
the of and notated, as type Miquel's species as belonging to Intsia bijuga (Colebr.)
Kuntze. There is also of this collection one duplicate in U, which had been annotated by G.J.H. Amshoff as belonging to Intsia.
Intsia tashiroi Hayata (1913) was based on a plant from S Ryukyu: Yaeyama (as
Yaemaya) Archipelago. I have not seen the type, but have studied the detailedoriginal description and examined well-prepared specimens, both in flower and in fruit, from
Ishigaki I., belonging to the same Archipelago, collected by S. Hatusima (1888, L), and Furuse (2837, 3712 & 4577, K, photos in L; 3727 & 3974, K). In the Index
Kewensis this species was erroneously listed as from Formosa (Taiwan); as far as known of found plants this genushave not yet been there in the wild (cf. Legumi- nosae in Fl. Taiwan, vol. 3, 1977: 148-421). Walker (1954) already rightly re-
this its distributionis further duced species to Intsia bijuga, so now known to extend north to southern Ryukyu (Fig. 1) (cf. De Wit, 1941: f. 1; Whitmore, 1972: f. 9).
2. Intsia palembanica Miq.
Intsia palembanica Miq., Fl. Ned. Ind., Suppl. (1860) 107, nom. nud.; (1861) 289; Prain, Sci.
Mem. Med. Off. Army India 12 (1901) 45; MeijerDrees, Bull. Jard. Bot. Buitenzorg III, 16
(1938) 92, f. lc; De Wit, Bull. Bot. Gard. Buitenzorg III, 17 (1941) 142, f. 1; Comer, Ways.
Trees ed. 2, 1 (1952) 396; Whitm., Tree Fl. Mai. 1 (1972) 262, f. 1, 2, 9; Meijer, Field Guide
Trees W. Mai. (1974) 185, f. 44; Cockb., Trees Sabah 1 (1976) 170, f. 36; Verde., Man. N.G. Ding Hou: Malesian Caesalpinioideae: Acrocarpus, Afzelia, Copaifera, Intsia 327
Legumes (1979) 93; K. & S.S. Larsen, R.Thailand 4 (1984) 126; Gunn, Techn. Bull. U.S.D.A.
1755 (1991) 258, f. A & D. — Afzelia palembanica Baker in Hook, f., R. Brit. Ind. 2 (1878)
275; K. Schum. & Lauterb., Nachtr. Fl. Deut. Schiitzgeb. (1905) 276; Ridley, R. Mai. Pen. 1
— (here chosen): (1922) 640; Burkill, Diet. 2 (1935) 1243, as I. bakeri Prain. Lectotype Teijs-
HB mann 4535 (U, photo in L), Sumatra (Lampong, near Kebang); syntypes: Teijsmann (all
no.) 3666, 3820, 3914 & 4315 (U, photos in L), Sumatra (Palembang & Lampong).
Intsia plurijuga Harms, Bot. Jahrb. 55 (1917) 54; Meijer Drees, Bull. Jard. Bot. Buitenzorg HI, 16
(1938) 95, f. Id; Kanehira & Hatusima, Bot. Mag. Tokyo 56 (1942) 363. — Syntypes; Leder- Schlechter K, in L), New Guinea. mann 7441 & 8652 (K, photos in L), 14219 (iso photo
90. Intsia acuminata auct. non Merr.: Verde., Man. N.G. Legumes (1979)
and Leaves 4-jugate, rarely associated with some 2-, 3-, or 5-jugate ones; petiole
Leaflets rachis (3.5—)9.5—17.5(—26) cm, both puberulous, glabrescent. ovate, ellip-
obovate tic, sometimesbroad-ovate or -elliptic, or suborbicular, rarely or lanceolate,
obtuse sometimes (3—)10—13.5(—18.5) by (3—)5—7.5(—10.5) cm; apex or rounded,
slightly notched, (shortly) acuminate; base obtuse, rounded, cuneate, rarely slightly
somewhat 6-10 truncate or subcordate, often asymmetric; nerves per side; petiolules
10 3-12 2-5 mm. Inflorescences up to c. cm long, pubescent, glabrescent; pedicels
3-4 lobes or mm. Hypanthium mm long. Calyx (broadly) elliptic, ovate, slightly
lamina obovate, 6—8(—12) by 3-6.5(-8) mm. Petal; 3-6(-10) by 3-6.5(-8) mm;
2-4 anthers 2 stami- claw 2-5 mm long. Fertile stamens; filaments cm; c. mm long;
5-6 2-3.5 Pods 5.5- nodes 4-10 mm. Ovary mm long; style cm. 15—19(—40) by
3-4.5 thick. 7(-9) cm. Seeds by 1.5-3 cm, 0.5-1.2 cm
- 45 m bole or sinuous, Field notes Tree up to (rarely more) high, straight slightly with buttresses up to 22(-30) m long, and up to l(-4) m d.b.h. Big trees often up
often rather to 7 m high, extending outwards 6 m, and 30 cm thick. Bark smooth,
red. Pods brown black not fissured, scaly. Petals pale yellow or white, pink, or or
dark brown. when ripe, smooth. Seeds reddish or
Distribution - India (Andamans), Burma, Thailand, and Malesia; throughout (ex-
1. cept Java ?). Fig.
Cultivated in Java (Hort. Bog., sub no. I-I-7 and I-C-10, from Sumatra).
of seasonal inundat- Habitat - Widespread, in coastal regions, edges swamps, on
low- ed lands, locally common to very common (e.g. in Japen I., New Guinea), in
land primary or older secondary forests, sometimes on sandstone and limestone
sea-level 850 alt. hills, from up to m
Ecology - Flowering from Jan.-April and Aug.-Nov. Fruiting from Feb.-Dec.
Uses - See under the genus.
- Vernacular names Sumatra: merbau abang, merbau sepang (M), tandoek (Palem-
merboh merbau toeh- bang), merbau asam, m. boei, m. gatja, m. insi, (Atjeh), tapah
vel (M), toeko (Nias). Malay Peninsula: ironwood & Malacca teak (English), merbau,
tandok Lesser Sunda Islands: rai m. ayer, m. kunyit, m. puteh, m. (M). (Wetar).
Brunei and Sarawak: merbau, m. ayer, m. bukit (Iban). Sabah: merbau (M), polo-
mok (Api-Api River). Kalimantan: alai, a. anglai (Dajak), djembai, kerek (Kutai),
ipil (M & Dajak), maharan (M). Celebes: ipi hout (Bone). Moluccas: dowora (Moro-
tai), dowora kome (Halmahera). New Guinea: bau (Atam), bauw, bouwa, sekka,
(Manikiong), bidjam, piam (Kebar), koboe, midek (Mooi), kwila (Madang & North-
ndirin ern Dist.), mer (Arfak), mewit, ngoeit (Karoon), (Tehid), pase (Asmat), piam
(Amberbakan), rompi (Japen), rong (Nemo), temmen (Tor). 328 BLUMEA VOL. 38, No. 2, 1994
Notes - There are two specimens from Sumatra collected by Teijsmann (s.n.) in
908.3-306 & annotated in Intsia L (under HLB 307), Miquel's handwriting as pa-
lembanica. These may be duplicates ofthe syntypes from U.
Podzorski (Palawan Bot. Exp. 1984, t. 2) published a photograph of the basal
named from the and part of a tree, as Intsia bijuga, Palawan, Philippines, indicated
that it is c. 4 m (four metres!) d.b.h. I have seen Ridsdale 366 (L), a specimen in
to the and thus extends its fruit, which belongs present species geographic range to
the Philippines.
Swollen branchlets occasionally have been observed, which may be caused by
ants (vide Brass & Versteegh 13542, L).
Meijer Drees (1938: 93) recorded that the trees flower usually after the fall of
leaves, so fruiting specimens have nearly always young fleshy leaves whereas sterile collections bear more or less leathery ones.
ACKNOWLEDGEMENTS
Curators Iam indebted to the Directors, Keepers, and of the following institutes forproviding facili- ties, assistance, and/or sending herbarium specimens on loan: BM, BO, K, KEP, L, SAN, SAR,
SING, and U.
My thanks are due to Dr L. Watson, Australian National University, for sending me a detailed
of the of in Malesia from the data bank. printout genera Caesalpinioideaeoccurring
I the late Dr R. Geesink for his cards of am grateful to my colleague, giving me bibliographic
Caesalpinioideaefor the Malesian flora.
is Dr W. Vink for the reviews the My appreciation due to my colleagues: preparing on regenera-
Peter Welzen for his useful tion and uses of the genus Intsia; Dr van giving me very computer pro-
M.C. grams of Herbarium Database Utilities; Dr P.W. I.eenhouts, Dr Roos, and Dr C. Kalkman for their critically going through the manuscript and helpful remarks; and Mr Jan van Os for pre-
the outline distribution paring map.
Identification list
1 = Acrocarpus fraxinifolius Arnott 4 = Copaifera palustris (Symington) De Wit
2 Leonard Intsia Kuntze = Afzelia javanica (Miq.) 5 = bijuga (Colebr.)
3 Intsia = Afzelia rhomboidea (Blanco) Vidal 6 = palembanica Miq.
Aet & Idjan 789: 5; 811: 6 — Agama 8856: 6 — Ahem 146: 3; 208: 5; 704: 3 — d'Alleizette
— 1951: 5 — Anang 381: 5 — Anderson, D. 987, 1201, 3703, 3739: 5 — Atasrip 79: 5 Atje
132: 5; 272: 6; 402: 5.
Backer 18747: 2; 33486: 5 — Balakrishnan, N.P. 3816: 5 — van Balgooy 2262: 6 — bb series
2226, 2700: 5; 3073: 6; 3918 (T), 4064: 5; 6124: 1; 6346: 2; 6380, 6381: 6; 6998: 5; 7003: 4;
7070: 6; 7187: 2; 7993: 6; 8365, 9142, 9353: 5; 9435: 6; 10125: 5; 13422, 15053, 15057: 6;
15063, 15081, 15370, 15968: 5; 16323, 16523, 16536, 16686: 6; 17624: 5; 19692: 6; 20412:
2; 20414, 21095, 21111: 6; 21476: 5; 22523: 6; 23038: 5; 23181, 23301: 6; 23828: 5; 24312:
6; 24454: 5; 24565: 6; 25012, 25074: 5; 25343: 6; 25489: 5; 25677, 25704: 6; 26003: 5;
26232, 27238, 28376: 6; 28734, 28764: 5; 28909: 6; 28998: 5; 29714: 2; 29800, 29845: 6;
30585, 30597: 5; 31066: 6; 31424, 31493, 31524: 5; 31600 (T): 2; 31663, 31731: 6; 31861,
31902: 5; 31991, 32171, 32206, 32375, 32460, 32580, 32581, 32583, 32588, 32589, 32590,
32591, 32592, 32593, 32594: 6; 32744: 2; 32763, 32764, 32765, 32767: 6; 32823, 32853,
33034: 5; 33253, 33280, 33378: 6; 33670, 33692, 33799, 33804: 5; 33826: 6; 33898: 5;
35079, 35161, 38376: 6 — Beccari PB 3915: 4 — Beguin 735: 5; 2205: 6; 2216, 2219: 5 — 329 Ding Hou: Malesian Caesalpinioideae: Acrocarpus, Afzelia, Copaifera, Intsia
40259: 6 — Brass 28550: Bemicr(?) 12034: 5 — BKF series 28699: 1; 40255: 5; 2975, 24293,
— 114: 5075: 7851: 6 5 — Brass & Versteegh 13542: 6 BRUN series 6; 5; 5102, 5642, 5643,
— — BS series 801: 5; 6982: 3; 17428, 26740: 5 BSIP series 109, 1081, 1934, 2689, 2924,
2951, 3158, 3930, 4236, 4716, 4996, 5330, 6240, 6546: 5; 6982: 3; 8643, 8883, 8998, 9420,
9670, 9728, 9791, 10376, 10467, 10672, 10813, 11384, 11662, 12344, 12619, 13103, 13639,
13700, 13804, 13927, 13975, 14037, 14333, 14424, 14543, 14596, 14741, 14945, 15361,
15485, 15706, 16397, 16508, 17192, 17549, 17601, 17671, 17951, 18248, 18458, 18949,
1088: 1160: 1325: 1428: 19499: 5 _BW series 47, 504, 693, 849, 5; 6; 1292, 1302, 5; 1388,
6; 1567: 5; 1598, 1746, 1895: 6; 1999: 5; 2079, 2087, 2092, 2096, 2201, 2288, 2292, 2618,
2830: 2631: 6; 2697, 2713, 2725: 5; 2742, 2786, 2823, 2824, 2825, 2826, 2828, 2829, 6;
3161: 2831, 2835, 2836: 5; 2838: 6; 2839, 2840, 2855: 5; 2916, 2970: 6; 5; 3244: 6; 3248,
4654: 3435, 3604, 3674, 3704, 3853, 3923: 5; 3976, 3980, 4247: 6; 4423: 5; 6; 4721, 4748,
4769: 5; 5606, 5748, 5875, 6308, 6744: 6; 6911, 7009: 5 ; 7206, 7239: 6; 7617: 5; 7667,
7707, 7845, 8088: 6; 8118: 5; 9005, 9083, 9098: 6; 9595, 9774, 9825, 9950: 5; 10015,
11000, 11084: 6; 11092, 11158: 5; 11628, 11765, 11852, 11998, 12340, 12907: 6; 14897,
15010: 5.
— Clemens Carr 11798: 5 — Cel series 111-47,111-97: 5 — Chew Wee Lek RSNH 303: 5 10556,
I-B-32: 10863: 5; 21289 (T): 3 — Cooper 3729: 1 — Cult, Hort. Bog. 1-1-2: 5; 1-1-7,1-C-10: 6;
2; I-C-16a: 5; I-L-30 & 31: 1; X-C-16a, XI-B-53: 5; XI-B-XV-11: 1 — Cuming 924: 3.
Dachlan 2371: 6 — Dumas 1562, 1566: 6.
Elmer 11144: 3; 12102, 12953: 5; 15317, 16803, 20878: 3 — Hndert 39E.1P.120, 39E.1P.366,
39E.1P.431, 39E.1P.473, 39E.1P.476, 39E.1P.480, 39E.1P.973, 39E.1P.974, E1351: 6 —
Eyma 2837: 5.
FB series 2046, 2570, 2591: 3; 4502: 5; 5450, 6642: 3; 8537, 9522, 10595, 10663, 12370, 12821:
5; 17773: 3; 17889: 5; 20780, 22607, 24605: 3; 25366: 5; 26605, 28152: 3 — F6nix 1148: 3
— — Flemmich EMS 32600 (T, n. v.): 4 Fosberg 24979, 25171, 26774, 26806, 26922, 32199,
34077, 34079, 34317, 34387: 5 — FRI Bogor 145T.1P.188: 3 — FRI series 1213, 4947: 6;
— 4577: 5. 5043: 5; 5811, 10590, 11090: 6; 11572: 5; 14514: 6 Furuse 2837, 3712, 3727, 3974,
Gjellerup 177: 5.
— Ham L3, B5: 5 — Hart, A.J. (NGF?) 5014: 5 — Hartley 0773, 9740: 5 Hatheway, W.H. 835:
— — 70903: 2 — 2855: 6 3439, 4364, 5 — Hatusima 1888: 5 Herb. Bogor Heyne Hoogland
— 4934: 5 —Hoogland & Craven 10298: 5 — Horsfield 'L 147' (T): 2; 'L 187' (T): 5 Hosaka,
E.Y. 2017, 3320: 5 — Hose 93 (T): 3 — Houter 57: 2 — Hunt, P.F. 2789: 5 — Hutchinson,
W.I. 205: 5 — Hyland 2823, 7460: 5.
Idjan & Mochtur 346: 6.
— 522: 5 — C. 617: 5 — Ja series 1068, 2544: 2; 2636: 5 — Jaheri 290: 5 Janowsky Jeffrey,
Jensen, H.J. 193, 354: 5.
— Keith 9883: 5 — KEP series 7908: 5; 14266: 6; 30426: 5; 32577: 4; 93491: 6 Kerr 9831: 1;
— 16584: 5 — Kingdon Ward 18416: 1 — KL series 3160: 5 — Koch, J.W.R. 634: 5 Koelz
24378, 27500, 32611: 1 — Koorders 3857, 3859, 3860, 3863: 1; 3869: 2; 3984, 3987, 3989,
3990: 5; 4151: 2; 4152, 4153, 4156, 13322, 13323: 2; 13805: 1; 14428, 14827, 14828,
15723: 2; 17554, 17555, 17556: 5; 17557: cf. 5; 17558: 6; 17559, 17560: 5; 27816: 1; 34000,
7762: 36913, 37020: 2 39264: 1 — Kornassi 211: 5 — Kostermans 5071: 6; 5; 9551, 10298:
— 6; 14061: 3; 18880: 2; 21884: 5; 22085: 6 — Kostermans et al. 362: 2 vanKregten 112: 5
— Krukoff 265: 2.
— Labohm 2012: 5 —LAE series 52447, 52662, 53793, 66055, 66747: 5; 70238: 6 Lam 3173: 5
— 6 — Loher 2201: 3 — — Leano 43: 3 Ledermann 7441 (T), 8652 (T): Liitjeharms 4201,
4227, 5147: 5.
Dist. 6 — Martati Mail 7057: 5 — Main & Aden 135: 6; 1604: 5 — Maingay (Herb.) 565 (K no.):
5 — Merrill 122: 1 — Maxwell 75: 5 — Meijer, W. 5636: 1; 9252: 1108 (T), 2584, 2594,
5 — C. 222: 5. — 128: Morrison, 2651: 3; Sp. Bl. 376 (T): 5; 862 (T): 3 Mondi
NGF series 1207, 1400, 1430, 2044: 5; 3789: 6; 4117, 15468, 20714, 22864, 26609, 26754,
— 32237, 32743, 36083, 37516, 42269: 5; 46555: 6 — van Niel 4084, 4555: 5 Noerkas 370: 6. 330 BLUMEA VOL. 38, No. 2, 1994
van Ooststroom 12902: 5.
— Parks, H.E. 20023, 20908: 5 Pleyte 147: 5 — Poilane 20290: 1.
Reynoso et al. 176, 329: 6 — Ridsdale 366: 6; 2175: 5 — van Royen 3282, 4096: 5; 5007: 6 —
Rutten 2019: 5.
S series 1210, 5219, 8261, 8565, 8852: 4; 11206: 6; 12268, 12942, 13111, 17257: 4; 25988: 3;
31884: 5 — SAN series A2679 (Kadir): 5; A 2724 (Mail),A 2973 (Melegrito): 3; 6711 (Puasa):
6; 8571 (Lupang): 5; 8572 (Lupang), 15308: 6; 17427: 4; 19871: 6; 24517: 3; 24929, 25757:
5; 25814: 6; 34583: 5; 35909, 36977, 37060: 3: 40477: 6; 42598, 44722, 47199: 3; 54987,
55710: 6; 56187, 56197, 57021: 3; 57252, 70224: 6; 75614, 82664: 3; 83716: 6; 82664,
88889, 89274, 93580: 3 — Saunders 196, 217, 531: 5 — Schlechter 14219 (T): 6 — Schodde
2747, 3082: 5 — Schodde (& Craven) 4545: 5 — SF series 37294 (cult.): 3; 39001: 5 — van
Slooten & Backer 35042: 5 — Smith, A.C. 9354: 5 — Stainton 6407: 1 — van Steenis 18240:
5 — Stone 3805, 4747: 5.
Takamine 251: 5 — Teijsmann HB 862 (T), HB 887 (T): 1; HB 2950: 5; HB 3666 (T), HB 3820
(T), HB 3914 (T), HB 4315 (T): 6; HB 4475: 5; 4535 (T): 6; HB 5026, 7800: 5 — Thakur Rup
Chand 4204: 1.
Uhl, F. 5953: 2 — Ursch 117: 5.
Verheijen 3384: 5 — Vidal 1282, 1287, 1297, 2635, 2670: 3 — Vieillard 386 (T): 5 — de Vogel
4254: 5 — Voyce 24: 5.
Wagapani, M. 2900: 6 — Wallich 5823, 5823A, 5823B: 5 — Waterhouse 293, 709-b, 938: 5 —
Wenzel 1692, 3122: 3 — Wight 2466 (T, n.v.), 845 (K Distr. no., T?): 1 — de Wilde & de
Wilde-Duyfjes 21134: 3 — Winit 169, 317: 1 — Wirawan 386: 5 — Worthington 1632, 1753,
2771: 1.